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Genetic Variability and Evidence of Founder Effect in Hemiodus Orthonops (Characiformes: Hemiodontidae) from the Upper Paraná River Basin, Brazil

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Genetic Variability and Evidence of Founder Effect in Hemiodus Orthonops (Characiformes: Hemiodontidae) from the Upper Paraná River Basin, Brazil Acta Scientiarum http://www.uem.br/acta ISSN printed: 1679-9283 ISSN on-line: 1807-863X Doi: 10.4025/actascibiolsci.v39i1.31793 Genetic variability and evidence of founder effect in Hemiodus orthonops (Characiformes: Hemiodontidae) from the upper Paraná River basin, Brazil Liégie Alher Marques1 and Erasmo Renesto1, 2* 1Centro de Ciências Agrárias, Universidade Estadual de Maringá, Maringá, Paraná, Brazil. 2Departamento de Biologia Celular e Genética, Universidade Estadual de Maringá, Av. Colombo, 5790, 87020-900, Maringá, Paraná, Brazil *Author for correspondence. E-mail: [email protected] ABSTRACT. Hemiodus orthonops is a small fish of the Hemiodontidae family, order Characiformes, with a maximum of 25 cm standard length. Until recently, H. orthonops was an endemic species from the Paraná- Paraguay basin and it was absent from the upper Paraná River basin. Since 2008, it has started to be collected in the upper Paraná River, representing up to 10% of catches. Two population samples of H. orthonops from two localities of the upper Parana River basin (Porto Camargo and Porto Figueira) were analyzed using the allozymes electrophoresis technique. Twenty-one enzymatic loci were detected. The population sample from Porto Camargo displayed a genetic variability (He = 0.1061) higher than that from Porto Figueira (He = 0.0580) and homozygote excess in both of them. The FST value (0.2081) indicated genetic structure. The excess of homozygotes in both samples was probably due to founder effect in the population. Keywords: Allozymes, fish, genetic distance, heterozygosity, protein polymorphism. Variabilidade genética e evidência do efeito do fundador em Hemiodus orthonops (Characiformes: Hemiodontidae) da bacia do alto rio Paraná, Brasil RESUMO. Hemiodus orthonops é um pequeno peixe da família Hemiodontidade da Ordem Characiformes com um comprimento padrão máximo de 25 cm. Até recentemente, H. orthonops estava ausente da bacia do alto rio Paraná. Desde 2008 ele passou a ser coletado na bacia do alto rio Paraná, representando até 10% das coletas. Duas amostras populacionais de H. orthonops provenientes de duas localidades da bacia do alto rio Paraná (Porto Camargo e Porto Figueira) foram analisadas pela técnica de eletroforese de aloenzimas. Vinte um loci enzimáticos foram detectados. A amostra proveniente de Porto Camargo revelou uma variabilidade genética (He = 0,1017) superior à amostra de Porto Figueira (He = 0,0558) e excesso de homozigotos em ambas as amostras. O valor de FST entre elas (0,2081) indica que há estruturação genética. O excesso de homozigotos nas duas amostras é provavelmente devido ao efeito do fundador. Palavras-chave: Aloenzimas, peixes, distância genética, heterozigosidade, polimorfismo proteico. Introduction provinces. As a consequence, 33 species of native fishes in the lower Paraná basin successfully Habitat alterations caused by water flow control induced by dams, for example, disrupt the structure colonized the upper Paraná River (Julio Junior, Dei of the aquatic biota and facilitate the establishment Tós, Agostinho, & Pavanelli 2009). In 2002, a of invasive species (Havel, Lee, & Zanden, 2005). channel for fish migration named “Canal de The installation of fish passages in dams, with the Piracema” was opened, allowing 28 of the 68 fish aim to preserve migratory species, may pose an species registered downstream of the dam to rise additional threat to the biota from upstream towards the Itaipu Reservoir (Agostinho, Gomes, stretches by increasing the dispersion of non-native Fernandez, & Suzuki, 2002). species. This may occur when the dam separates Hemiodus orthonops is a small fish of the distinct native faunas along the watershed, as Hemiodontidae family, order Characiformes, with a observed in the Paraná River (Agostinho et al., maximum of 25 cm standard length (Froese & 2015). In the case of Paraná River, the Itaipu Pauly, 2015), described by Eigenmann & Kennedy Reservoir, filled in 1982, covered the Sete Quedas (1903) from specimens collected in the Paraguay Falls, which separated distinct ichthyofauna River, near the town of Asuncion. Acta Scientiarum. Biological Sciences Maringá, v. 39, n. 1, p. 53-58, Jan.-Mar., 2017 54 Marques and Renesto Until recently, H. orthonops was an endemic some of them it was even higher. Bossdorf et al., species from the Paraná-Paraguay basin and it was (2005) revealed that 69% of invasive plants had the absent from the upper Paraná River basin same or greater genetic diversity than native (Agostinho et al., 2015). Since 2008, it has started to populations. A similar study of 29 terrestrial and be collected in the upper Paraná River, representing aquatic animal species showed that introduced up to 10% of catches (Agostinho et al., 2015). species have preserved 80% of the genetic variability In the Paraná River population studied by existent in native populations (Wares, Hughes, & Agostinho et al. (2015), the maximum standard Grosberg, 2005). length observed for H. orthonops was 26 cm for males Thus, in this study we aimed to estimate the and 29 cm for females; individuals become sexually genetic variability of Hemiodus orthonops using the mature at one year of age when they reach 15.2 cm allozymes electrophoresis technique and compare it of length for males and 19.4 cm for females, to other species from Paraná River basin, which was releasing a maximum amount of 79,653 oocytes also studied with allozymes electrophoretic with 0.79 mm diameter each and feeding primarily technique. Furthermore, we verified the presence of on detritus and algae. a founder effect in the invasive population of this According to Agostinho et al. (2015), the success migratory species in the Paraná River. of H. orthonops at colonizing the upper part of Paraná River was due to dispersal ability, favorable Material and methods environmental conditions, the ability to exploit Hemiodus orthonops individuals were collected in highly available food resources, early maturation and the upper Parana River basin, area of Porto Camargo high body growth rate. (23°21’27.87”S, 53°44'54.14"W), municipality of Invasive species are predicted to suffer from Icaraíma and in Porto Figueira (23°23'43.93"S, reductions in genetic diversity when only a few 53°48'47.79"W), municipality of Alto Paraíso (Figure individuals from the source population colonize 1) since March, 2012 to April, 2013. new habitats, thus undergoing the founder effect and reducing adaptive potential (Mayr, 1963; Allendorf & Luikart, 2007). The founder effect on genetic variability of experimental populations of mosquito fish were measured by the allozymes electrophoretic technique by Leberg (1992), who estimated the values of proportion of polymorphic loci, number of alleles per locus and heterozygosity. Compared to source populations, Leberg found that 23 populations had a smaller He and 15 had an equal or greater He. On average, all three indexes of genetic variability decreased as the effective size of the founding population decreased. Roman and Darling (2007) reviewed data of 43 introduced aquatic species and found that only 16 (37%) showed clear evidence of significant loss of genetic diversity relative to native populations, while in 27 species there was no alteration. Dlugosch and Parker (2008) analyzed data from 80 species in the literature covering 18 plants, 2 fungi, and 60 animals (including 7 birds, 6 reptiles, 8 fish, 3 amphibians, 8 mammals, 13 insects, 4 crustaceans, 6 mollusks, 3 annelids, 1 cnidarian, and Figure 1. Map showing the places of capture of Hemiodus 1 tunicate) and showed that, in introduced orthonops: Porto Camargo (23°21’27.87”S, 53°44'54.14"W) and Porto Figueira (23°23'43.93"S, 53°48'47.79"W), State of Paraná, populations, there was an average reduction of Brazil. 15.5% in allelic richness and a decrease of 18.7% in average heterozygosity compared to the original Fifteen specimens were captured in the first population. mentioned municipality, and 13 in the second one. However, other studies have shown that many Voucher specimens were deposited in the collection invasive species did not lose the variability and in of Nupelia (Núcleo de Pesquisas em Limnologia, Acta Scientiarum. Biological Sciences Maringá, v. 39, n. 1, p. 53-58, Jan.-Mar., 2017 Genetic variability of Hemiodus orthonops 55 Ictiologia e Aquicultura) of the Universidade Estadual de calculated the mean of 74 species distributed in 12 Maringá NUP 10610. All specimens were frozen and families (49 Loricariidae, three Callichthyidae, five stored at low temperature (-20° C) until the Pimelodidae, one Cynodontidae, one Erythrinidae, extraction of enzymes. The enzymes were extracted six Anostomidae, one Prochilodontidae, one from samples of muscle, liver and heart tissue, using Auchenipeteridae, three Gymnotidae, one buffer Tris-HCl, 0.02 M, pH 7.5. The samples were Serrasalmidae, three Characidae and one Cichlidae). centrifuged in a refrigerated centrifuge at 4° C and 16,434.6 g. The protein extract was applied on corn Results starch gel (Val, Schwantes, Schwantes, & De Luca, In the Porto Camargo sample (PC), 27 alleles 1981) Penetrose 50 at a concentration of 17%, with distributed in 21 loci were detected in H. orthonops, small filter paper strips (4 mm x 8 mm) Whatman 3 six loci of which were polymorphic (Adh, G6pdh, MM® soaked with the samples. Subsequently, the Ldh-1, Est-1, Est-2, Est-3); in Porto Figueira sample gel was submitted to a continuous horizontal (PF), 25 alleles were detected in 21 loci, of which electrophoresis under refrigeration. Three buffer three were polymorphic (Adh, Est-1, Est-3). solutions were used: 0.135 M Tris/Citric Acid 0.043 Table 1 illustrates the allele frequencies of each M, pH 5.9 (TC), diluted 15 times in the preparation locus of each sample. The Aspartate enzyme Amino of the gel; 0.1 M Tris/Maleic Acid 0.1 M/EDTA Transferase (AAT) expressed two loci: Aat-1, which 0.004M/Magnesium chloride 0.01M, pH 7.4 (TEM) codifies cathodic enzymes and Aat-2 that codifies in tank and diluted 10 times in the gel, and Tris anodyne enzymes.
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